The occurrence of high molecular weight humic material in recent organic-rich sediment from the Namibian shelf

The molecular size distribution of humic substances has been investigated in two recent sediment samples of different ages taken from a single core sample collected off Walvis Bay, south-west Africa. Extractable humic acids were found to be by far the major form of organic carbon in both sediments, the near-surface younger sample containing predominantly high molecular weight (>100 000 molecular weight) humic acids and little fulvic acids whilst the deeper, older sample contained relatively less humic acid and relatively more fulvic acid of a range of molecular weights. A significant age difference was found between the >300 000 molecular weight and the <30 000 molecular weight fractions of the near-surface sample, the lower molecular weight fraction being older than the higher molecular weight fraction.The data suggest that in this rapidly accumulating, organic-rich sediment the first step en route from the planktonic matter to the humic complexes is direct and rapid incorporation of the biogenic material into high molecular weight humic acids.     

Petroleum hydrocarbons in sediments of Southampton water estuary

The sediments of Southampton Water were analysed for petroleum hydrocarbons to determine the fate of petrochemical refinery waste in the estuary. Much of the petroleum hydrocarbons appears to be removed by adsorption onto estuarine sediments close to source. Surface sediment concentrations range from 3·1mg/g dry weight near a refinery to 0·5 mg/g dry weight at locations distant from the refinery.The absence of any change with depth in total hydrocarbon concentrations from some sediment cores and the presence of distinct ‘oil’ horizons in others suggests that mixing of sediments in the area is very random. Sedimentation rates in areas where a distinct oil horizon was observed ranged from 1 to 3cm a year.     

Dynamics of carbohydrates in the Norwegian Sea inferred from monthly profiles collected during 3 years at 66°N, 2°E

Water samples were collected monthly for 3 years at 66°N, 2°E in the Norwegian Sea, 250 nautical miles off the Norwegian coast. Concentrations of mono- and polysaccharides were measured with the 2,4,6-tripyridyl-s-triazine (TPTZ) spectroscopic method. Total dissolved carbohydrates varied from 3.4 to 28.2 μM C of all samples and the ratio of carbohydrate to dissolved organic C (DOC) varied from an average of 14% at 0–25 m depth to 11% at 800–2000 m depth. This indicates that dissolved carbohydrates were a significant constituent of DOC in the Norwegian Sea. Polysaccharides varied from 0.4 to 21.5 μM C and monosaccharides from 0.7 to 11.7 μM C at all depths. The level of monosaccharides was relatively constant at 2.8–3.2 μM C below the euphotic zone, whereas polysaccharides showed more varying concentrations. Dissolved carbohydrates accumulated during the productive season, reaching maximum concentrations during summer although interannual differences were observed. A significant positive correlation between Chl a and soluble carbohydrate was found in one growing season with nutrient analyses. Average values for total carbohydrates were highest in the surface – 0 to 25 m – with 13.3 μM C and decreased to 8.4 μM C at 800–2000 m depth. The ratio of monosaccharides to polysaccharides exhibited a marked seasonal variation, increased from January to a maximum in June of 1.1, and declined to 0.5 in July.     

A study of the formation of high molecular weight compounds during the decomposition of a field diatom population

This work has investigated the possible formation of humic and fulvic acids, particularly high molecular weight moieties, in degrading diatom debris. The diatom debris was collected, freshly sedimented, from a well characterized natural field diatom population and allowed to degrade under conditions similar to those found at the sediment-water interface of many marine, organic-rich sediments.Samples of the detritus and overlying water were taken regularly over a period of 4 months and analysed for the presence of humic compounds. In the case of the detrital material, a sequential series of extractions were used. Molecular weight fractionation of the extracted humic material was carried out using Diaflow Ultrafiltration Membranes and two different fractionation methods have been compared. It appears that the choice of method may have an important effect on the determined distribution of the humic material, particularly when humic acid concentrations are low.High molecular weight (> 300 000- > 100 000) humic and fulvic acids were found in all the detrital samples, the content increasing with the period of decomposition. No evidence of any humic material was found in extracts of the living diatoms, the conclusion being that this material had been formed in situ during the degradation of the diatom debris. Most (65–90%) of the humic material was extracted from the detritus using a mild 0·05 m NaOH extraction at room temperature. Fulvic acids comprised approximately 90% of the total 0·05 m NaOH extraction, being composed mainly of either low molecular weight compounds (40–50% < 10000) or high molecular weight compounds (38–48% > 100000). Levels of humic acids were much lower and included a wide range of molecular weight fractions.The relevance of these findings to the possible origin of humic material in certain marine sediments is discussed.     

Molecular weight and trace metal distributions in fulvic and humic acid fractions of coastal marine sediments

Molecular weight and trace metal distributions of fulvic and humic acid fractions of marine sediments from the Seto Inland Sea were investigated by using a gel filtration technique in combination with atomic absorption and emission spectrometries. A binary molecular weight distribution was found both in the fulvic acid fraction and in the humic acid fraction. The fulvic acid fractions with molecular weights of less than 1.5×10³ and of 5×10³?10⁴ accounted for 38–57 % and 25–41 % of the total fulvic acids, respectively, and the humic acid fractions with molecular weights of 10²?10⁴ and of over 2×10⁵ accounted for 58–73 % and 16–27 % of the total humic acids, respectively. The components with molecular weights of over 1.5×10³ contained 54 %, on average, of the Fe, Zn and Cu in the fulvic acid fraction, and the components with molecular weights of over 10⁴ contained 58 %, on average, of these metals in the humic acid fraction.     

Cadmium accumulation by the blue crab, callinectes sapidus: Involvement of hemocyanin and characterization of cadmium-binding proteins

The accumulation of cadmium from seawater by the blue crab, Callinectes sapidus, was studied as a function of metal concentration and exposure time, with special emphasis on cadmium-binding proteins. Cadmium was found, in decreasing order of magnitude, in gills, digestive gland and hemolymph. When exposed to 0·5 ppm cadmium for 2–24h, virtually all of the cadmium in the cytosolic fraction of the gill was associated with a low molecular weight (LMW) cadmium-binding protein (MW 8000). However, after 48h of exposure only 50% of the cadmium in the cytosol was bound to this protein. The rest wasfound to be associated with proteins of a molecular weight of 300 000 and 60 000. This pattern of cadmium distribution did not change over a 12-day depuration period. Similar results were obtained upon exposure to 0·1 ppm cadmium. The pattern of cadmium accumulation in the cytosolic fraction of the digestive gland was in marked contrast to that observed for the gill. Initially, the cadmium was distributed over three low molecular weight fractions. During depuration the distribution of cadmium changed and all of the metal became bound to a low molecular weight protein (MW 9000). The cadmium concentrations in the gill and digestive gland remained essentially constant during depuration (12 days). The LMW cadmium-binding proteins were purified by a combination of gelpermeation and ion-exchange chromatography. Their molecular weight, spectral properties and amino acid composition are characteristic of the vertebrate metallothioneins. During exposure to cadmium the metal rapidly appeared in the hemolymph, mainly associated with hemocyanin. During depuration cadmium was transferred from the hemolymph to the digestive gland, demonstrating that hemocyanin acts as a carrier in trace metal transport.     

A giant piston-corer

A new large piston-coring device utilizing well-casing pipe 14 cm (o.d.), 11 cm (i.d.) has obtained cores over 20 m in length in water depths of 100 to 5000 m. Recovery ratios range from 0·76 to 0·87. Penetration varies between 24 and 29 m with a total core weight of 3200–4000 kg and 2 to 3 m of free fall.     

Studies in seawater and lake water on interactions of trace metals with humic substances isolated from marine and estuarine sediments: I. Characterisation of humic substances

Two humic acids, HAL and HAN, isolated from marine sediments (Adriatic and Norwegian Sea) and humic (HAM) and fulvic (FAM) acids isolated from an estuarine sediment (Mahakam, Borneo) were characterized.The following characteristics: carbon content, hydrophobicity, degree of condensation and the content of aliphatic groups, are most pronounced for HAN, less pronounced for HAL and the least pronounced for HAM. The abundance of trace metals in the humic acids decreases in the following order: Al > Cu > Zn > Pb > Cd.Fulvic acid (FAM) has a lower carbon and a higher oxygen content than humic acids. It is hydrophilic and hardly any aliphatic groups were recorded. At pH 7 the predominant fraction of FAM has the apparent molecular weight of 20 000 daltons while the humic acids are of higher molecular weight.     

Carbohydrates in humic and fulvic acids from Hiroshima Bay sediments

A core of 75 cm length from Hiroshima Bay, Seto Inland Sea in Japan, has been analyzed for the carbohydrate content of the humic and fulvic acids. These carbohydrates were found by gel filtration to be in the high molecular weight range. Carbohydrates were more abundant in fulvic acids than in humic acids. A comparison of carbohydrates in humic acids with those in fulvic acids showed that the former did not undergo diagenetic loss any faster than the latter. Identification of most alditol acetate in monosaccharides is based on the comparison of gas chromatographic retention indices and mass spectrometric fragmentation with those of authentic standard compounds. However, the compositional differences between carbohydrates in humic and fulvic acids were greater in the top section of 0–20 cm than in the 50–70 cm section. The processes which produce the humic-acid carbohydrates are operative either prior to or shortly after deposition.     

Chemical studies of dissolved carbohydrates in seawater

Charcoal column chromatography and a dialysis system using a Visking tube were used for the concentration and separation of dissolved carbohydrates in seawater. Recoveries of mono-, oligo-, and polysaccharides were 46–74, 61–89 and 70%, respectively, relative to the total amount of sugars initially submitted to analysis. The combined charcoal column chromatography and dialysis system was used to separate dissolved carbohydrates in seawater samples from Mikawa Bay into mono-, oligo- and polysaccharide fractions. The polysaccharides are separated into two fractions with molecular weights of <4,000 and >4,000 using Sephadex G-25 column chromatography. Monosaccharides, oligosaccharides, polysaccharides with M.W.<4,000 and polysaccharides with M.W. >4,000 accounted for 7–9, 15–26, 7–13 and 20–33% of total dissolved carbohydrates, respectively. However, 32–38% of dissolved carbohydrates were lost during the analytical procedures. Glucose, galactose, mannose, xylose, arabinose, ribose, fucose, and rhamnose were found upon acid hydrolysis of the oligo- and polysaccharide fractions, while fucose and rhamnose were not present in the monosaccharide fraction. Possible sources of the carbohydrates in these fractions are discussed with reference to the monosaccharide composition of cellular carbohydrate in marine algae.     

Copper in the resurrection fjord, Alaska

Copper concentrations have been measured in more than 200 samples collected from an Alaskan fjord and continental shelf and slope regions in the northwestern Gulf of Alaska. Concentrations were lowest (2·1 nmol kg⁻¹) at depths of 400–1000 m in the continental slope waters of the Gulf of Alaska. Copper increased systematically with decreasing salinities shoreward to concentrations >30 nmol kg⁻¹ in fjord surface waters during summer months of high freshwater runoff. Copper concentrations increased with depth at an inner fjord station where deep basin waters have restricted circulation, and these data together with surface (<5 cm) pore water copper concentrations (mean=122 nmol kg⁻¹) about an order of magnitude higher than bottom water copper concentrations are indicative of a flux of copper across the sediment-seawater interface. This latter was estimated at 32±12 nmol cm⁻² annually, and represented less than 20% of the annual input to fjord surface water (228–411 nmol cm⁻²) added during summer months. Mass balances in bottom waters indicate a vigorous recycling of copper with a residence time estimated at 21±11 days. Most copper that is remobilized in surface sediments is returned to bottom waters and little (3%) is removed by subsequent diagenetic reaction in the buried sediments. However, an estimate of copper accumulating in anoxic fjord sediments was comparable with copper added to fjord surface waters suggesting that input-removal reactions rather than internal cycling controls copper geochemistry in this estuary.     

Influence of Particle Mixing on Vertical Profiles of Chlorophyll a and Bacterial Biomass in Sediments of the German Bight, Oyster Ground and Dogger Bank (North Sea)

In May and September 1999 11 stations were sampled in the southern and central North Sea, located in the German Bight, eastern Oyster Ground and Dogger Bank. The study focused on the influence of particle mixing on transport of chlorophyll a to deeper sediment layers and vertical bacterial distribution (max. DEPTH=10 cm). The sampling stations were chosen to reflect a gradient in environmental conditions in the North Sea. The sampling stations differed in respect to redox potential (eH up to −243 mV in the German Bight and up to 274 mV in the offshore regions), silt content (up to 54% in the German Bight and 0·34% at the northern Dogger Bank) and different proportion of fresh organic material on total organic matter content (C/N ratios ranging from 9·27 in the German Bight up to 1·72 in the offshore sediments). Although bacterial densities (8·55×10⁹ g⁻¹in the German Bight up to 0·35×10⁹ g⁻¹in offshore sediments) were significantly correlated to chlorophyll a content in the sediment (P<0·01), inconsistencies in the temporal pattern of both variables in the surficial sediment layer suggested, that the dynamics of bacterial densities is generally controlled by food supply but also by other variables. The chlorophyll a content in the surficial sediments of the German Bight (up to 1·84 μg g⁻¹) was significantly higher than in the Oyster Ground (up to 0·58 μg g⁻¹) and the Dogger Bank area (up to 0·68 μg g⁻¹). With increasing chlorophyll a input to the benthic realm a subsequent enhanced burial of this compound into deeper sediment layers was expected either by biological (bioturbation) or by physical sediment mixing. However, the vertical profile of chlorophyll a decreased steeply in the sediments of the German Bight. Contrary, subsurface peaks were measured in the offshore areas. It was concluded from these results, that the vertical distribution of organic matter in sediments is less limited by the quantitative input from the water column but concomitant with particle mixing itself. The extent and possible mechanisms of particle mixing in the different study areas in relation to specific environmental factors is discussed.     

Controls on the organic chemical composition of settling particles in the Northeast Atlantic Ocean

The organic matter of sinking particulate material collected in the Northeast Atlantic Ocean (ca. 49°N, 16°W) was investigated in order to determine temporal and depth-related variability in its composition. Three sediment traps were deployed at nominal depths of 1000 m (below the permanent thermocline), 3000 m (representing the deep-water fluxes) and at 4700 m, about 100 m above the seafloor (just above the benthic boundary layer). The samples span a 28-month sampling period from October 1995 until February 1998, each sample representing a period of between 7 and 28 days.Total organic carbon and total nitrogen contents decrease with depth, as did the absolute concentrations of most biochemicals measured in this study, such as intact proteins and individual lipids. However, concentrations of proteins relative to total organic carbon and total nitrogen did not show any significant change with depth, implying that they are not being rapidly degraded and so may provide an important supply of nitrogen to the benthos. Fluxes of protein, TN and TOC are significantly correlated at all depths.Lipid compositions vary temporally. During periods of high flux, particularly in the summer, the lipids are richer in ‘labile components’, namely unsaturated fatty acids and low molecular weight alcohols. During periods of low flux other compounds, such as sterols, steroidal ketones and a trisnorhopan-21-one are more abundant. One sample, taken close to the seafloor, was highly enriched in lipids, sterols and fatty acids in particular; this may represent detritus derived from bottom-dwelling invertebrates.     

Relationships between macroalgal biomass and nutrient concentrations in a hypertrophic area of the Venice Lagoon

Macroalgae biomass and concentrations of nitrogen, phosphorus and chlorophyll a were determined weekly or biweekly in water and sediments, during the spring-summer of 1985 in a hypertrophic area of the lagoon of Venice. Remarkable biomass production (up to 286 g m⁻² day⁻¹, wet weight), was interrupted during three periods of anoxia, when macroalgal decomposition (rate: up to 1000 g m⁻² day⁻¹) released extraordinary amounts of nutrients. Depending on the macroalgae distribution in the water column, the nutrients released in water varied from 3·3 to 19·1 μg-at litre⁻¹ for total inorganic nitrogen and from 1·8 to 2·7 μg-at litre⁻¹ for reactive phosphorus. Most nutrients, however, accumulated in the surficial sediment (up to 0·640 and to 3·06 mg g⁻¹ for P and N respectively) redoubling the amounts already stored under aerobic conditions, Phytoplankton, systematically below 5 mg m⁻³ as Chl. a, sharply increased up to 100 mg m⁻³ only after the release of nutrients in water by anaerobic macroalgal decomposition. During the algal growth periods, the N:P atomic ratio in water decreased to 0·7, suggesting that nitrogen is a growth-limiting factor. This ratio for surficial sediment was between 6·6 and 13·1, similar to that of macroalgae (8·6–12·0).     

Benthic microalgal biomass in sediments of Onslow Bay, North Carolina

Sediment samples were collected at stations along cross-shelf transects in Onslow Bay, North Carolina, during two cruises in 1984 and 1985. Station depths ranged from 11 to 285 m. Sediment chlorophyll a concentrations ranged from 0·06 to 1·87 μg g⁻¹ sediment (mean, 0·55), or 2·6–62·0 mg m². Areal sediment chlorophyll a exceeded water column chlorophyll a a at 16 of 17 stations, especially at inshore and mid-shelf stations. Sediment ATP concentrations ranged from 0 to 0·67 μg g⁻¹ sediment (mean, 0·28). Values for both biomass indicators were lowest in the depth range including the shelf break (50–99 m). Organic carbon contents of the sediments were uniformly low across the shelf, averaging 0·159% by weight. Photography of the sediments revealed extensive patches of microalgae on the sediment surface.Our data suggest that viable benthic microalgae occur across the North Carolina continental shelf. The distribution of benthic macroflora on the North Carolina shelf indicates that sufficient light and nutrients are available to support primary production out to the shelf break. Frequent storm-induced perturbations do not favour settling of phytoplankton, an alternative explanation for the presence of microalgal pigments in the sediments. Therefore, we propose that a distinct, productive benthic microflora exists across the North Carolina continental shelf.     

Settleable and Non-Settleable Suspended Sediments in the Ogeechee River Estuary, Georgia, U.S.A.

Suspended particle dynamics were investigated in the Ogeechee River Estuary during neap tide in July 1996. Samples were operationally separated into ‘ truly suspended ’ (settling velocity <0·006 cm s⁻¹) and ‘ settleable ’ (settling velocity >0·006 cm s⁻¹) fractions over the course of a tidal cycle to determine whether these two fractions were comprised of particles with differing biological and chemical characteristics. Total suspended sediment, organic carbon and nitrogen, chlorophyll a and phaeopigment concentrations were measured in each fraction, as well as rates of bacterial hydrolytic enzyme activity [β-1,4-glucosidase (βGase) and β-xylosidase (βXase)]. The majority of the suspended sediment (by weight) was in the truly suspended fraction; all measured parameters were largely associated with this fraction as well. When compared to the settleable material, the truly suspended material was significantly higher in % POC (5·7±0·6 vs. 3·9±1·8), % chlorophyll (0·07±0·02 vs. 0·03±0·01), % phaeopigment (0·030±0·006 vs. 0·018±0·012), and weight-specific maximal uptake rates (V_maxper mg suspended sediment) of both enzymes (1·8±0·4 vs. 0·7± 0·2 nmol mg⁻¹ h⁻¹βGase and 1·1±0·3vs . 0·3±0·2 nmol mg⁻¹ h⁻¹βXase), providing clear evidence for a qualitative distinction between the two fractions. These results are interpreted to mean that the more organic-rich, biologically active material associated with the suspended fraction is likely to have a different fate in this Estuary, as ‘ truly suspended ’ sediments will be readily transported whereas ‘ settleable ’ sediments will settle and be resuspended with each tide. These types of qualitative differences should be incorporated into models of particle dynamics in estuaries.     

Phosphorus and metals bound to organic matter in coastal sediments - An investigation of complexes of P,Cu, Zn,Fe, Mn,Ni, Co and Ti by inductively coupled plasma-atomic emission spectrometry with sephadex gel chromatography

Phosphorus and metals bound to organic matter were separated from coastal sediments of Harima Sound in Seto Inland Sea, Japan by extraction with NaOH and fractionated by Sephadex G-25 chromatography. Phosphorus and metals were determined in the eluates by a multi-channel, inductively coupled plasma-atomic emission spectrometer. Phosphorus and Cu, Zn, Fe, Mn, Ni, Cr, Co and Ti bound to organic matter with high molecular weights (OMHMW) (MW ? 5000) were found to be present in the sediments, but no Mo or V were found. The technique provides minimum estimates of the amounts of P and metals bound to organic matter. These organic complexes show surface enrichment in a sediment core (0–20 cm) and their contents decrease with depth. Also, the amounts of eighteen elements, namely: P, Fe, Mn, Zn, Cu, Si, Al, Ti, Pb, Co, Ni, Cr, Mo, V, Na, K, Ca and Mg, in H₂O, ammonium acetate at pH 7 and 5, hydrogen peroxide, hydroxylamine hydrochloride and hydrogen fluoric acid soluble fractions have been determined with a selective chemical leaching technique for the ²¹⁰Pb-dated sediment core sample. Considerable amounts of P (6–19%) and Cu (5–21%) were associated with organic matter, in contrast to other metals such as Fe, Mn, Zn, Ni, Cr, Co and Ti which were associated with sulfide and silicate.     

Biogeochemistry of organic matter in the Laurentian Trough, II. Bulk composition of the sediments and relative reactivity of major components during early diagenesis

The organic carbon of 280–320 m deep Laurentian Trough sediments at landward and seaward sites (13–24 mgN/g) consisted of carbohydrates (15–22%), hydrolysable amino acids (7–13%), lipids (1–5%), labile proteins (0.3–1%) and a non-characterized fraction (62–74%). Amino acids, proteins and uncharacterized compounds accounted for 21–43, 0.9–4 and 51–78%, respectively, of total nitrogen (1.2–2.2 mgN/g). A clear reactivity trend (pheopigments ? lipids > proteins > amino acids ≈ nitrogen > carbon > carbohydrates) was deduced from the concentration decreases between settling particles and surficial sediments. This was confirmed by one-year inventories in the top cm, burial rates at 35 cm depth, and one-G model calculations. Lipids were a dominant substrate near the sediment-water interface whereas carbohydrates and amino acids constituted the principal energy sources deeper in the sediment. In the porewaters, DOC levels were low (2–6 mg/l) in the top 4 cm, indicating rapid removal (i.e. consumption, irrigation, diffusion), and increased with depth (8–12 mg/l), reflecting the buildup of refractory products. There were also clear compositional changes of DOC with depth. Geographical differences in water column fluxes were recorded in the sediments. The organic contents and ratios were higher at the landward site due to higher rates of sedimentation, bioturbation and terrestrial and total organic inputs. At the seaward station, the lower rates of these processes and stronger marine influence resulted in lower ratios and a more complete decay of organic matter within the top 35 cm sediments.     

Variability in in situ shear strength of gassy muds

Torque measurements are made by divers with a vane apparatus to a sediment depth of 136 cm in the high-porosity, gassy sediments of Eckernförde Bay. Corrected shear strength values calculated from torque measurements are quite variable in the Eckernförde Bay sediments, varying from less than 0.5 kPa in the top 10 cm of sediment to 4–5 kPa at 136 cm sediment depth. Variability increases markedly below 60 cm sediment depth, probably because of the presence of methane gas bubbles within the sediment fabric.     

Distribution of nitrogenous nutrients and denitrifiers strains in estuarine sediment profiles of the Tanshui River, northern Taiwan

Chemical profiles of both oxidized (nitrate and sulfate) and reduced (ammonium, sulfide, acid-volatile sulfide [AVS], and pyrite) materials and the corresponding distribution of denitrifier microbial communities were measured at low tide in sediments at Guandu in the estuary of the Tanshui River, northern Taiwan in August 2002. Denitrifier strains were isolated for physiological and phylogenic analyses. Based on the distribution of nitrogenous compounds and denitrifier abundances, the vertical profile of Guandu sediments could be separated into four layers: a mixed layer (the top 1 cm of depth, respectively containing 0.82–2.37 and 535.9–475.0 μM of nitrate and ammonium), a nitrate-concentrated layer (1–5 cm in depth, 2.37–0.53 and 475.0–1192.1 μM, respectively), a denitrifier-aggregation layer (5–7 cm in depth, 0.53–0.72 and 1192.1–1430.1 μM, respectively), and an ammonium-enriched layer (7–12 cm in depth, 0.72–0.78 and 1430.1–2196.6 μM, respectively). Denitrifier strains were detected in all layers except for the mixed layer. A variety of metabolic processes by these strains may occur in different layers. Bacillus jeotgali-, Bacillus sphaericus-, and Bacillus firmus-related strains isolated from the nitrate-concentrated layer may be involved in the nitrification-denitrification coupling process due to the relatively low nitrate concentrations (maximum = 2.37 μM), and may contribute to denitrification not nitrification. Bacillus bataviensis- and B. jeotgali-related strains isolated from the denitrifier-aggregation layer comprised the predominant denitrifier population (3.64 × 10⁴ cells/g of denitrifier abundance). They possess the ability of dissimilatory nitrate reduction to ammonium (DNRA). Bacillus jeotgali-related strains and two newly identified strains of GD0705 and GD0706 isolated from the ammonium-enriched layer possibly use fermentative processes as the main metabolic pathway instead of denitrification when nitrate is scarce, and this further supports the high ammonium concentrations (up to 2.20 mM) found in the Guandu sediments. In addition, spore formation also enhances the chance of survival of these strains in the face with such a nitrate-deficient environment.